Flora of the Canadian Arctic Archipelago
Poaceae, Grass family.
Published in Can. J. Bot. 62: 2454. 1984.
Type: Greenland, Jameson Land, Gurreholm, 14 VIII. 1958, Holmen 807. Holotype: C. Isotype: O!
Synonymy. F. vivipara L. subsp. glabra Frederiksen, Nordic J. Bot. 1: 288. 1985.
Festuca ovina L. var. vivipara L., pro parte. Sp. Pl. ed. 2, 1: 108. 1762.
Festuca brevifolia R. Br. var. arctica Saint-Yves subv. genuina f. vivipara Saint-Yves, Candollea 2: 253–254. 1925. Type: Greenland. Arakamtchetchem, Herb U.S.N. Pacif. Expl. Exp. 1853–56, leg. Wright. Holotype: P.
Festuca brachyphylla f. vivipara Skvotscov. in Tolmatchew, Fl. Arct. URSS, 222–223. 1964, nom. nud.
Festuca ?viviparoidea Krajina, Biota N. Amer. 2: 342. 1980, nom. nud.
Vegetative morphology. Plants 11–28 cm high (bluish grey-green); perennial herbs; caespitose; sometimes vegetatively proliferating in inflorescences. Only fibrous roots present. Ground level or underground stems absent. Aerial stems erect. Leaves mainly basal; alternate; marcescent. Prophylls 10–20 mm long; with hairy veins; with pronounced keels. Petioles absent. Sheaths present; with the margins fused only in the lower part; glabrous, or with trichomes (that are minute); hirsute; sheath collars present. Ligules present; 0.2–0.5 mm long; membranous; hairy (ciliate at the apex); transversely oblong. Ligule apices truncate (and higher on sides than middle); erose and cleft. Leaves grass-like. Blades 30–120 mm long, 0.35–0.75 mm wide (when folded, with limited sclerenchyma strands seen in leaf cross sections that are similar to those of F. brachyphylla and unlike those of F. ovina), spreading, folded in bud (with margins inrolled and sometimes overlapping), linear, involute, veins parallel, midvein similar in size to other veins in the leaf (usually 3 larger veins, 2–4 smaller veins). Blade adaxial surface glabrous or hairy. Blade abaxial surface hairy.
Reproductive morphology. Flowering stems circular or oval in cross section. Flowering stems with leaves; culm nodes not exposed, or becoming exposed; culm nodes number visible 0–1. Flag leaf sheaths not inflated. Inflorescences with bulbils; paniculate; dense; linear; 1–3 cm long; 5–15 mm wide. Inflorescences main axis scabrous. Number of inflorescence branches at lowest node 0–1 (smaller inflorescences have a single spikelet per rachis node). Inflorescence primary branches 0.2–0.6 mm long; scabrous; with appressed secondary branches. Spikelets disarticulating above the glumes; lanceolate; 7–15 mm long (or longer, including the vegetative proliferation); 2–3 mm wide. Florets per spikelet 2–3. Two glumes present (unequal.). First glume 0.7–0.8 × the length of the second glume; 0.3–0.45 × spikelet length; 3–4 mm long; lanceolate; glabrous, or with trichomes (hairs at the apex only); margins ciliate; veins 1; apex acuminate. Second glume 0.4–0.9 × as long as the spikelet; almost as long as, or longer than, the lowest floret; 3.6–6 mm long. Second glume lanceolate. Second glume glabrous, or with trichomes (scabrous at apex only); veins 3. Rachilla not pronounced between the florets; terminating in a vestigial floret (or terminating in a vegetatively proliferating floret); internode 0.1–0.3 mm long. Lemma lanceolate; 3.6–6 mm long; keeled (more pronounced as lemma structure becomes leaf-like in vegetative proliferation); surface dull (deep purple); surface sparsely scabrous; surface with trichomes on and between the veins; veins 5; apex acuminate; apex entire; apex glabrous; awnless, or awned (often awnless in proliferating spikelets). Awn arising from the tip. Awn 0.1–0.3 mm long. Palea vestigial, or absent (rarely formed). Flowers bilaterally symmetrical (zygomorphic); bisexual (or replaced by vegetative proliferation). Perianth represented by lodicules (absent in proliferating spikelets). Stamens present, or absent (sometimes aborted anthers present); 0–3. Anthers splitting longitudinally (if applicable). Anthers usually not developed. Ovary superior. Ovaries ovaries rarely if ever developed. Styles absent. Ovules per ovary 1 (rarely applicable). Fruit a caryopsis (sometimes vegetatively proliferating).
Chromosome information. 2n (4x) = 28. Tzvelev's draft for Elven et al (2003);
2n (7x)= 49. ?Flovik (1938, Siberia); Frederiksen (1981, Greenland);
2n (8x) = 56. Johnson and Packer (1968, northwestern Alaska, as F. vivipara); Frederiksen (1981, northern Alaska);
2n (9x) = about 63. Holmen (1964, Alaska, three counts).
Ploidy levels recorded 4x/7x/8x/9x.
Taxon as an environmental indicator. The phenomenon of vegetative proliferation in grasses has been observed to be a response of plants to environmental stresses such as over-watering or over-fertilising in greenhouse conditions. If further collections of this taxon are made in the High Arctic, very thorough habitat notes would be relevant. It is possible that the collections to date are vegetatively proliferating plants of either F. brachyphylla or F. baffinensis, growing under environmentally stressful conditions on Northern Ellesmere Island, but there has been limited material on which to base this conclusion. Festuca rubra. proliferates in cold, damp sites along the North American east coast, and it is possible that the colder, harsher conditions on Northern Ellesmere trigger the proliferation found in plants called F. viviparoidea.
Ecology and habitat. Substrates: hummocks, tundra, slopes; imperfectly drained moist areas; gravel (and turf). Occurring in tundra turf, gravel, and moist herbmats. Reported from Northern Ellesmere Island. The limited records are of plants with spikelets that are beginning to vegetatively proliferate. Except in this character they cannot be distinguished from specimens of either Festuca baffinensis or F. brachyphylla.
North American distribution. Alaska, Yukon, continental Northwest Territories (?), Nunavut Islands (?). Arctic islands: Ellesmere.
Northern hemisphere distribution. Circumpolar (with gaps). Svalbard Franz Joseph Land, Polar Ural Novaya Zemlya, Taimyr Severnaya Zemlya, West Chukotka, Wrangel Island, South Chukotka, East Chukotka, West Alaska, North Alaska Yukon, Ellesmere Land Peary Land, East Greenland.
General notes. Porsild (1964) mentions F. vivipara (L.) Sm., noting that the Canadian specimens are similar to F. baffinensis and F. brachyphylla, but the panicle is always proliferous. Frederiksen (1981) stated that "according to a handwritten note by Holmen, the chromosome number given for Plantae Vasculares Groenlandicae Exsiccatae nr. 22 belongs to the above-mentioned holotype. The number 2n = 56 was published (Holmen and Laegaard 1960), but re-examination shows 2n = 49 to be correct, and as far as I know, no other numbers have been known from Greenland... 2n = 56 was found in material from Alaska and Wrangel Island." This species has very often been connected with F. brachyphylla, and morphologically it looks very much like this species. In the few cases where anthers were observed, they were about 2 mm. As F. brachyphylla is very distinct because of its very short anthers, rarely if ever exceeding 1.2 mm (Frederiksen 1977), and as F. brachyphylla has the leaf sheaths partly split while they are always open in F. vivipara, no close connection seems to exist between these two taxa. On the other hand F. vivipara subsp. glabra seems isolated from the two other subspecies in nearly all examined characters; thus it possibly has another origin.
A specimen at CAN, which was annotated by Signe Frederiksen in 1987, was collected on Ellesmere Island [Lower Dumbell Lake, the water supply lake for Alert, growing in moist herbmat C.R. Harrington 201, August 18, 1959]. An extensive search, in 1992, to find the voucher for the mapped record from Ellesmere Island, Judge Daily Promitory (Frederiksen 1981), was unsuccessful. The specimen from the Canadian Arctic may be an F. baffinensis ? F. brachyphylla hybrid. The plants have very young inflorescences, with few definitive characteristics. Several attempts to obtain fresh material with which to do isozyme experiments have failed.
Festuca vivipara s.s. was interpreted by Alexeev as an offspring from F. ovina, alone, or in combination with other species. The triploid chromosome number was interpreted by Frederiksen (1981) as a result of hybridisation between F. ovina and F. vivipara subsp. vivipara. She stated, however, that the triploid plants are morphologically indistinguishable from the tetraploids. They occur independently outside the range of F. ovina and form large populations (Salvesen 1986). Triploids may have evolved several times as the hybrid F. vivipara ×ovina, but most of the southern Norwegian triploids seem to constitute a distinct alpine taxon. Tetraploid F. vivipara cannot easily be derived from any ancestral form occurring in Scandinavia today. From morphological and geographical evidence, it therefore seems reasonable to include both triploids and tetraploids (and probably also higher polyploids) in F. vivipara subsp. vivipara (Elven, in Elven et al. 2003).
Alexeev (1995) interpreted F. viviparoidea as a viviparous offspring of F. brachyphylla. Festuca viviparoidea is based on subsp. glabra and the same type. Frederiksen (1981) explicitly stated that she found F. vivipara subsp. glabra to belong to the 'ovina' group, not the 'brachyphylla' group. Pavlick (1984) reported the anthers, when present, to be of 'ovina'-group length. It is therefore very improbable that Frederiksen's subsp. glabra (and by implication F. viviparoidea) are anything else than a viviparous offspring from the F. ovina group rather than the F. brachyphylla group. Morphologically, the material from northern Siberia (for example, Taimyr), northern Alaska and northwestern Canada differs from the Atlantic F. vivipara and might have an origin in the F. brachyphylla group. It is unlikely that the name F. viviparoidea is correctly applied for them.
Elven et al. (2003) noted that the vegetatively proliferating plants found in the Canadian Arctic Archipelago have affinities with the F. brachyphylla complex.
Viviparous plants occur fairly regularly in 'continental' northwestern North America, in British Mountains, and northern Alaska where they form populations, have their own ecology, and behave like a taxon. There may be a gap in the distribution between this area and Greenland, making the identification of the northwestern American plants with F. vivipara subsp. glabra (and thereby with F. viviparoidea) problematic.
Several attempts to obtain living material of suitable plants from Northern Ellesmere Island have failed. Isozyme analyses of such plants would have been conclusive if they were merely vegetatively proliferating specimens of either F. baffinensis or F. brachyphylla. The above information is included to draw attention to collections from the Arctic Islands where further study is required to determine the correct taxonomic status.
Illustrations. • Specimen of plant. Festuca plant with vegetativey proliferating inflorescences. Alaska, Delta River, Darling Creek. R. Elven and H.H. Grundt. Herb. Oslo. • Close-up of spikelets. Spikelets with glumes and above them vegetatively proliferating florets. Yukon Territory, Ivvavik Natl. Park, Buckland Hills. R. Elven. 2103/99. Voucher at O. • Arctic Island Distribution.
This publication is available on the internet (posted May 2011) and on CD-ROM (published in 2007). These versions are identical in content, except that the errata page for CD-ROM is accessible on the main index page of the web version.
Recommended citation for the web-based version of this publication: Aiken, S.G., Dallwitz, M.J., Consaul, L.L., McJannet, C.L., Boles, R.L., Argus, G.W., Gillett, J.M., Scott, P.J., Elven, R., LeBlanc, M.C., Gillespie, L.J., Brysting, A.K., Solstad, H., and Harris, J.G. 2007. Flora of the Canadian Arctic Archipelago: Descriptions, Illustrations, Identification, and Information Retrieval. NRC Research Press, National Research Council of Canada, Ottawa. http://nature.ca/aaflora/data, accessed on DATE.
Recommended citation for the CD-ROM version of this publication: Aiken, S.G., Dallwitz, M.J., Consaul, L.L., McJannet, C.L., Boles, R.L., Argus, G.W., Gillett, J.M., Scott, P.J., Elven, R., LeBlanc, M.C., Gillespie, L.J., Brysting, A.K., Solstad, H., and Harris, J.G. 2007. Flora of the Canadian Arctic Archipelago: Descriptions, Illustrations, Identification, and Information Retrieval. [CD-ROM] NRC Research Press, National Research Council of Canada, Ottawa..